Records of the Western Australian Museum 20: 333-344 (2001).

Copepods from ground waters of Western Australia, V. Phyllopodopsyllus wellsi sp. novo (Crustacea: Copepoda: ) with a key to world species

2 T. Karanovicl, G.L. Pesce and W.F. Humphreys3

1 Via Brescia 3, 84092 Bellizzi (SA), Italy 2 Dipartimento di Scienze Ambientali, University of L'Aquila, Via Vetoio, 67100 L'Aquila, Italy 3 Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia

Abstract - Phyllopodopsyllus wellsi sp. novo is described from anchialine ground waters of the Cape Range karst area in north-western Australia and has clear stygomorphies, being colourless and lacking the nauplius eye. The genus is for the first time recorded from Australia, as well as for the first time from outside strictly marine habitats. A key to all the species of the genus Phyllopodopsyllus T. ScoU, 1906 is presented.

INTRODUCTION absence, and dimensions of an unguiform Arid north-western Australia is proving to projection). Coull (1973) presented a very detailed contain a diverse stygofauna inhabiting both survey of the genus Phyllopodopsyllus, and compiled ancient freshwater systems (Bradbury and an instructive table listing the most salient Williams, 1997; Poore and Humphreys, 1999a; morphological characteristics. His key included 33 Watts and Humphreys, in press) as well as species and subspecies. Finally Kunz (1984) anchialine waters (Humphreys, in press). Recently, reviewed the genus, and tabulated setal formulae of a number of stygal cyclopoid have been the second, third, and fourth swimming legs in described from the ground waters of Western female, as well as the antennulae segmentation and Australia in both near-coastal (Pesce et al., 1996a, shape. He subdivided the genus into nine species 1996b; Pesce and De Laurentiis, 1996) and ancient groups, based on these characteristics: 1 - bradyi, 2 ­ continental regions (De Laurentiis et al., 1999). The furciger, 3 - aegyptieus, 4 - borutzkyi, 5 - pauli, 6 ­ present paper concerns an harpacticoid opistoeeratus, 7 - mossmani, 8 - xenus, and 9 ­ from anchialine ground waters of north-western longipalpatus group. Piers (1995) doubted the Australia, belonging to a genus previously recorded "naturalness of these groups", but proposed only in strictly marine habitats. nothing new, placing his new species (P. According to Bodin (1997), the tetragonicipitid yucatanensis) within one of Kunz's groups. genus Phyllopodopsyllus T. Scott, 1906 is, one of the Subsequent to the last review of the genus (Kunz, most speciose marine harpacticoid genera, together 1984) 13 new species and subspecies have been with Halectinosoma Lang, 1944 and Stenhelia Boeck, described: P. alatus Fiers, 1986; P. crenulatus Wells 1865, and marine harpacticoid genus with the and Rao, 1987; P. graeilipes Wells and Rao, 1987; P. largest number of new described species in the last stigmosus Wells and Rao, 1987; P. tenuis Wells and fifty years. Lang (1948) initialy divided the genus Rao, 1987; P. galapagoensis Mielke, 1989; P. kunzi Phyllopodopsyllus into two genera (Phyllopodopsyllus Mielke, 1989; P. aneylus Mielke, 1992; P. earinatus and Paraphyllopodopsyllus), based upon the shape of Mielke, 1992; P. mossmani ehiloensis Mielke, 1992; P. the second segment of the antennula and setation of hartmannorum Kunz, 1995; P. pallaresae Kunz, 1995; second and third swimming legs in females. Many and P. yueatanensis Fiers, 1995. In addition Wells authors followed this revision, and Vervoort (1964) and Rao (1987) described the male of P. aegypticus provided an useful key to species of the genus Nicholls, 1944 and Kunz (1995) described the male Phyllopodopsyllus, although not for Paraphyllo­ of P. xenus (Kunz, 1951). Kunz (1984) however did podopsyIIus. After several new species were not mention three species already described: P. described possessing intermediate characteristics, laticauda Por, 1964; P. medius Por, 1964; and P. Lang (1965) synonymized Paraphyllopodopsyllus bahamensis Geddes, 1968. Even though some of the with Phyllopodopsyllus and compiled an excellent species above are synonyms, the number of species key to 21 species them recognized. He subdivided in the genus PhyllopodopsyIIus has increased the genus into three morphological groups based considerably, and a key for their identification is on the second antennular segment (i.e. presence or therefore included in this paper. During an 334 T. Karanovic, G.L. Pesce, W.F. Humphreys

investigation of the copepod fauna in Western Permanent mounts were made in commercial Australia a new species of the genus polyvinyl-Iactophenol and in Faure's medium. Phyllopodopsyllus was identified. This new species is Dissected specimens were drawn at magnification herein described as P. wellsi sp. novo of 630x and 1000x with drawing tube mounted on a Leica DMLS microscope, with C-plan achromatic objectives. All material is deposited in the Western HABITAT AND ASSOCIATED FAUNA Australian Museum (WAM) and BES denotes field The copepods were sampled from ground water numbers from the Invertebrate Biogeography and in karst terrain, accessed by temporary bores drilled Ecology group. Abbreviations used in the key to for a seismic survey. The bores were located on a species are: Fu - furcal rami; A1- antennula; P1-P4 wave-cut limestone terrace overlain by Quaternary - first to fourth swimming leg; P5 - fifth leg; Enp ­ deposits on the north-western tip of the Cape Range endopodite; Exp - exopodite; Enp2P3 - second peninsula, Western Australia. Bore A5 lies one endopodite segment of the third swimming leg. kilometre south-east of Babjarrimannos at an altitude of a 12 m ASL and ca. 1000 m from the seashore; the water table approximates sea level SYSTEMATICS 3 and had a salinity of 24.7 g/dm • Bore A7 is within one kilometre of Vlaming Head at an altitude of a Family Tetragonicipitidae Lang, 1944 10 ASL and ca. 350 m from the seashore; the water 3 Genus Phyllopodopsyllus T. Scott, 1906 is about sea level and had a salinity of 25.7 g/dm • Ground water on the eastern side of the peninsula, Phyllopodopsyllus wellsi sp. novo where it has been recorded, tipically tracks the Figures 1-19 marine tides but exhibits a temporal and amplitude lag (Humphreys et al., 1999). The waters are Material Examined typically anchialine (Stock et al., 1986; Sket, 1996; Humphreys, 1999b) and exhibit a marked physico­ Holotype ~ chemical stratification (Humphreys, 1999b). Fauna (WAM C24456), Cape Range, AB7, Western sampled within the same boreholes as Phyllo­ Australia, Australia, 21°49'S, 114°06'E, 11 podopsyllus wellsi sp. novo were all stygal species and November 1995, leg. RD. Brooks (BES 4676.1). included the blind gudgeon Milyeringa veritas Whitley (Pisces: Eleotridae), calanoid copepods and Allotype melitid amphipods. These form part of a more ;:) (WAM C244457), Cape Range, AB7, Western widespread and diverse stygofauna inhabiting both Australia, Australia, 21°49'S, 114°06'E, 11 fresh and anchialine waters of the Cape Range November 1995, leg. RD. Brooks (BES 4676.1). peninsula, with elements being found on Barrow Island 170 km to the north-east (Humphreys, 1999b; Paratypes in press; Humphreys et al., 1999). The following Australia: Western Australia: 14 ;:), 11 ~ (9 cyclopoid copepods are known from the area: ovigerous) (WAM 24458 - C24465, on slides, Apocyc1ops dengizicus (Lepechkine), Halicyc10ps C24466 in alcohol) Cape Range, AB7, 21°49'S, longifurcatus Pesce, De Laurentiis and Humphreys, 114°06'E, 11 November 1995, leg. RD. Brooks (BES Metacyc10ps mortoni Pesce, De Laurentiis and 4676.2). Humphreys, Microcyc1ops varicans G.O. Sars, and Diacyc10ps humphreysi Pesce and De Laurentiis. In Other material addition, the wider anchialine fauna contains Australia: Western Australia: 3 ;:), 5 ~ (4 numerous widely disjunct taxa, many of which ovigerous) (WAM C24467, in alcohol), Cape Range, occupy a "Tethyan" track, such as Lasinectes AB7, 21°49'S, 114°06'E, 5 November 1995, leg. RD. (Remipedia), Halosbaena (Thermosbaenacea), Brooks (BES 4673). Haptolana (Isopoda), Liagoceradocus (Amphipoda), 1 ~ (WAM C24468, on slide), Cape Range, AB5, Danie1opolina (Ostracoda), as well as misophrioid 21°50'S, 114°05'E, 4 November 1995, leg. R.D. and calanoid copepods (Yager and Humphreys, Brooks (BES 4666). 1996; Poore and Humphreys, 1992; Bruce and Humphreys, 1993; Bradbury and Williams, 1996; Description Danielopol et al., in press). Female (holotype) Habitus elongated, cylindrical. Body colourless, MATERIAL AND METHODS and nauplius eye absent. Cuticle densely furnished The sites were sampled using baited traps and with minute pits, especially on furcal rami and haul-nets from 'uphole' bores drilled to locate antennula (Figures 1 and 2). Prosome comprising recording equipment for a seismic survey. cephalothorax, incorporating first pedigerous Phyllopodopsylllls wellsi sr. novo from ground waters of Western Australia 335

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3

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Figures 1-4 Phyllopodopsylllls wellsi sp. nov., holotype (female): 1 antennula and rostrum; 2 - abdomen, dorsal view; 3 - maxilliped; 4 - mandibula. Scales = 0.1 mm. 336 T. Karanovic, G.L. Pesce, W.F. Humphreys somite, and 3 free pedigerous somites. Surface of armed with 1 lateral and 2 apical (1 smooth and 1 dorsal shield covering cephalothorax with many plumose) setae (Figure 4). Both rami I-segmented. sensillae, as well as tergites of 3 free pedigerous Maxillula with 13 setae on precoxa, 5 setae on somites. Hind margins of somites smooth. Rostrum coxa, 8 on basis, 4 on endopodite and 3 setae on almost quadrate, with single pair of sensillae, and exopodite. with row of very fine spinules on distal margin Maxilla with 10 seta on syncoxa (2 setae on each (Figure 1). Urosome comprising fifth pedigerous of proximal 2 endites; 3 setae on each of distal 2 somite, genital double-somite (representing fused endites), 4 on basis and 6 setae altogether on 2­ genital and first abdominal somite), and 3 free segmented endopodite. abdominal somites. Genital double-somite almost as Maxilliped with unornamented syncoxa, which long as wide (Figures 6 and 9). Subdivision line armed with 3 plumose setae (Figure 3). Basis with 1 visible dorsally, furnished with 6 sensillae, while seta and row of long spinules on inner margin. laterally and ventrally marked by rigid internal Endopodite I-segmented, armed with strong sclerotized ridge. One ventral and one dorsal pair recurved apical claw and slender smooth subapical of sensillae presented posteriorly on genital double­ seta (Figure 3). somite, as well as on first free abdominal somite All swimming legs with 3-segmented exopodites (Figure 2); second free abdominal somite lacking and 2-segmented endopodites (Figures 5, 7, 8 and sensillae. Their hind margins smooth, too. Anal 11). Spine and setal formula on exopodites and somite (last abdominal somite) ornamented only endopodites from first to fourth swimming legs with pair of sensillae dorsally. Anal operculum (legend: inner / outer spine or seta; inner / terminal convex, not reaching beyond limit of anal somite, / outer): with many marginal spines (Figure 2). Exopodite Endopodite Furcal rami divergent, about 3.8 times longer than Segments 1 2 3 1 2 wide, with complete armature (2 lateral, 1 dorsal First leg 0/1 0/1 0/2/2 1/0 0/2/0 and 3 apical setae), and without dorsal chitinous Second leg 1/1 0/1 1/2/2 % 0/3/0 ridge. Their inner margins with row of fine hairs, Third leg 1/1 0/1 1/2/2 1/0 0/2/0 especially long in distal part (Figure 2). Dorsal seta Fourth leg 1/1 0/1 3/2/2 1/0 0/2/0 attached at distal sixth of furcal length, well Couplers (intercoxal sclerites) of all swimming developed, while lateral setae very short. Outer legs with concave distal margin and without surface apical seta somewhat moved to ventral side, as long ornamentation. Coxa of third and fourth swimming as inner apical seta. Middle apical seta legs unornamented, while coxa of second leg extraordinary strong, slightly curved at end, and bearing row of hairs on inner-distal corner and coxa long as other two apical setae. of first leg ornamented with row of spinules on Antennula 8-segmented, with 1 slender aesthetasc outer margin. Basis of first and second swimming on apical segment,l very long aesthetasc on fourth legs with row of hairs on inner margin, while those segment (more than twice longer than 4 distal of third and fourth legs without ornamentation. segments taken together), and with setal formula as Basis of first swimming leg bearing 1 strong spine follows: 1.8.5.2.1.2.4.7 (Figure 1). All setae smooth. on inner margin (attached at first third of segment's Second segment with very large, posteriorly length). All swimming legs with smooth epipodite directed, unguiform process, and lacking anterior setae, on outer margin of basis. Apical and outer process. First segment about 3 times as long as elements on distal exopodite segment of fourth wide. swimming leg reduced, setiform (Figure 5). Antenna with almost completely reduced coxa, 1­ Fifth lag typically foliaceous, with completely segmented basis and exopodite, and 2-segmented fused basoendopodite and exopodite, but with clear endopodite (Figure 12). Inner margin of basis chitinous suture marking original separation furnished with long spinules. Exopodite with 3 (Figure 13). Epipodite seta well developed, and elements, of which outer one basally fused with between it and proximal margin of basis 1 row of segment. First endopodite segment only with few slender spinules exists. Former basoendopodite small spinules on outer margin. Second endopodite armed with 4 slender setae. Former exopodite segment with 2 lateral spines and 6 apical setae, as segment armed with 6 setae, of which only well as with many spinules along outer margin innermost one plumose. Fifth seta on that segment, (Figure 12). from inner margin, characteristically curved (Figure Mandibula with cutting edge of coxa with row of 13). Fifth leg cavity without eggs. teeth of different thickness and with single lateral Sixth leg fused with genital double-somite, seta (Figure 4). Basis large, bearing 3 setae on inner bearing 2 smooth setae, of which outer one about margin (1 smooth and 2 plumose), and furnished 1.8 times longer than inner (Figure 6). Genital field with 2 transverse rows of long slender spinules. present in distal half of first genital somite, having Endopodite twice longer than exopodite, armed short copulatory ductus connecting wide with 2 lateral and 4 apical smooth setae. Exopodite copulatory pore and small paired seminal Phyllopodopsyllus wellsi sp. novo from ground waters of Western Australia 337

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Figures 5-9 Phyllopodopsylllls wellsi sp. nov., holotype (female): 5 - fourth swimming leg; 6 genital double-somite, ventral view; 7 - second swimming leg; 8 - third swimming leg; 9 genital double-somite, dorsal view. Scales 0.1 mm. 338 T. Karanovic, G.t. Pesce, W.F. Humphreys

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Figures 10-13 Phyllopodopsyllus wellsi sp. nov.; 10, allotype (male); 11-13, holotype (female): 10 - abdomen, lateral view; 11 - first swimming leg; 12 antenna; 13 - fifth leg. Scales = 0.1 mm. Pllyllopodopsyllus wellsi sp. novo from ground waters of Western Australia 339

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Figures 14-19 Phyl/opodopsyl/us wel/si sp. nov., allotype (male): 14 - antennula and rostrum; 15 third swimming leg; 16 fourth swimming leg; 17 second swimming leg; 18 fifth leg; 19 - genital somite with sixth leg, lateral view. Scales = 0.1 mm. 340 T. Karanovic, G.L. Pesce, W.F. Humphreys receptacles. Small, tongue-like, spermatophore distal endopodite segment of the third swimming attached to genital pore (Figure 6). leg can have either 2 or 3 apical setae. In specimens with 3 setae, the pattern is very similar to that of the Male (allotype) distal endopodite segment of the second swimming Body similar to female in general appearance, leg in females (Figure 7). Sometimes left and right slightly more slender. Urosome comprising fifth third legs of the same specimen can bear different pedigerous somite, genital somite, and 4 free number of setae on that segment. No such abdominal somites. Genital somite about 1.5 times variability has been noted in the number of setae broader than long. All somites with smooth hind and spines of the other swimming legs. The margins, and with different number of sensillae. exopodite of the male fifth leg with or without very Furcal rami similar to female, but outer and small sharp extension on outer-distal corner middle apical setae much longer (Figure 10). Outer (between second and third setae, from outer side). apical seta more than twice longer than inner apical Moreover, sometimes one leg displays such one, while middle apical seta almost as long as extension, while opposite leg does not in same abdomen. . The outer seta on the female sixth leg can Antennula strongly geniculate, with almost be more or less plumose. Anal somite in female completely fused fourth and fifth segments (only 1 usually with 1 row of very small, equal, spinules at cuticular suture remained between them), and with base of furcal rami ventrally, but sometimes with a completely fused sixth and seventh segments few larger spinules in the middle of that row. (Figure 14). First and second, as well as ultimate, Holotype female is without eggs, but ovigerous segments very similar to female. females always with only two large eggs in the fifth Antenna, mandibula, maxillula, maxilla, leg cavity. The eggs are with very thin chorion. maxilliped, first swimming leg, exopodite of second (Figure 17) and third swimming legs (Figure 15) Etymology also very similar to those of female. The species is named in honour of Dr J.B.J. Wells, Outermost apical seta on distal endopodite Victoria University of Wellington, New Zealand. segment of second swimming leg very strong and basally fused with segment (Figure 17). Innermost apical seta on that segment short and smooth. DISCUSSION Endopodite of fourth swimming leg similar to Phyllopodopsyllus wellsi sp. novo differs from all female, except for inner apical seta on distal other species in the genus by its characteristic furcal segment being much stronger, curved, and smooth rami shape in the female, and especially by the (Figure 16). Distal exopodite segment of fourth middle apical seta, which is very short, stout, and swimming leg with only 5 appendages (2 inner, 1 slightly and characteristically curved distally. apical and 2 outer). Basoendopodite of the fifth leg in male with Basoendopodite of fifth leg with 3 strong, innermost seta the longest is also a rare characteristic plumose and spiniform endopodite setae in the genus Phyllopodopsyllus. Only three other (innermost seta longest), and with very long and species possess a similar basoendopodite: P. xenus smooth epipodite seta (Figure 18). Exopodite (Kunz, 1951); P. paraxenus Coull, 1970; and P. hermani subquadrangular, somewhat longer than broad, and Coull, 1969. P. xenus has indeed only two setae on armed with 5 elements (of which innermost spine that segment (Kunz, 1995), and differs from the new extraordinary strong and longest). Outer-distal species also by many other characters such as corner of exopodite distinctly produced, but antennula segmentation, swimming legs setation, without sharp extension. furcal rami shape, etc. P. paraxenus has 9-segmented Sixth leg consisting of small triangular chitinous antennula without unguiform process on second plate bearing 3 setae (Figure 19). Middle seta about segment, possesses different setation of swimming 1.6 times longer than inner one, and 2.8 times longer legs, and a different furcal rami shape (Coull, 1970). than outermost seta. Also, the middle seta on fifth leg basoendopodite in male of that species is extremely short. P. hermani Variability differs from the new species by many characteristics Thirty-six specimens (18 males and 18 females) of (Coull, 1969). Regarding the shape of the fourth Phyllopodopsyllus wellsi sp. nov., from 3 different swimming leg endopodite in female (1 seta on basal localities, were found and examined. Eleven and 2 setae on distal segment) P. wellsi is similar specimens were completely dissected and mounted with the four following species: P. minutus Lang, on slides. Body length, measured from the tip of 1948; P. bahamensis Geddes, 1968; P. paraxenus Coull, rostrum to the posterior margin of furcal rami, 1970; and P. tenuis Wells and Rao, 1987. They all ranges from 0.41 mm to 0.523 mm (0.455 mm differ from the new species by their swimming leg average) in males, and from 0.47 mm to 0.592 mm setation and shape of the furcal rami, whereas P. (0.527 mm average) in females. In both sexes the bahamensis and P. paraxenus also differ by the Phyllopodopsyllus wellsi sp. novo from ground waters of Western Australia 341

antennula segmentation (Lang, 1948; Geddes, 1968; Exp3P2 and Exp3P3 with 5 setae/spines .. Coull, 1970; Wells and Rao, 1987). P. wellsi does not ...... P. wellsi n.sp. fit well within any of nine species group created by Kunz (1984), which maybe confirms suspicions 14. Enp1P3 without seta 15 about its naturalness expressed by Fiers (1995), but Enp1P3 with inner seta 21 the philogenetical grouping of the species is outside 15. Exp2P4 without inner seta . the scope of this paper. Because we do not want to ...... P. medius Por, 1964 create a new group for this species, we provide a key to all species and subspecies in the genus Exp2P4 with inner seta 16 Phyllopodopsyllus. In that key have used mainly 16. Enp2P2 and Enp2P3 with 2 setae . female morphological characteristics, unless stated ...... P. hartmannorum Kunz, 1995 otherwise. Enp2P2 and Enp2P3 with 3 setae 17 17. Enp1P1 without inner seta .. Key to species and subspecies of the genus ...... P. punctatus Kitazima, 1981 Phyllopodopsyllus Enp1P1 with inner seta 18 1. EnpP4 I-segmented 2 EnpP4 2-segmented 3 18. Exp3P2 with 4 setae/spines ...... P. laticauda Por, 1964 2. Enp1P2 and Enp1P3 with inner seta .. Exp3P2 with 5 setae/spines 19 ...... P. geddesi Kunz, 1984 Enp1P2 and Enp1P3 unarmed . 19. A18-segmented P. bermudae Lang, 1948 ...... P. opisthoceratus Geddes, 1968 Al 9-segmented 20 3. Enp1P4 unarmed 4 20. Enp1P1 almost as long as ExpP1 .. Enp1P4 with inner seta 8 ...... P. angolensis Kunz, 1984 Enp1P1 about 1.5 times longer than ExpP1 ...... 4. Al 9-segmented P. xenus (Kunz, 1951) ...... P. aegypticus Nicholls, 1944 Al 8-segmented 5 21. Enp1P2 without seta 22 5. ExpP1 2-segmented . Enp1P2 with inner seta 32 ...... P. biarticulatus (Wells, 1967) ExpP1 3-segmented 6 22. Exp2P4 without inner seta 23 Exp2P4 with inner seta 24 6. Enp2P4 with 3 setae ...... P. longipalpalpatus hawaiiensis Kunz, 1984 23. Al 9-segmented P. borutzkyi Lang, 1965 Enp2P4 with 2 setae 7 A18-segmented P. simplex Kitazima, 1981 7. Fu with dorso-Iateral cuticular process .. 24. Exp3P3 with 5 setae/spines . .... P. longipalpatus madagascarensis Kunz, 1984 .. P. langi Kunz, 1975 Fu without that process . Exp3P3 with 6 setae/spines 25 P. longipalpatus longipalpatus (Chappuis, 1954) 25. Al 9-segmented 26 8. Enp2P4 with only 1 seta . Al 8-segmented 28 ...... P. hermani Coull, 1969 26. Fu about 7 times as long as wide .. Enp2P4 with more than 1 seta 9 ...... P. longicaudatus A. Scott, 1909 9. Enp2P4 with 2 setae 10 Fu less than 2.5 times as long as wide 27 Enp2P4 with 3 setae 14 27. Fu with large proximal inner bulge . 10. A19-segmented 11 ...... P. bradyi (T. Scott, 1892) Al 8-segmented 12 Fu without proximal inner bulge ...... P. pallaresae Kunz, 1995 11. Al with unguiform process ...... P. bahamensis Geddes, 1968 28. Al with additional sharp process on the anterior distal corner of second segment .... 29 Al without unguiform process ...... P. paraxenus Coull, 1970 Al without additional sharp process Gust one unguiform, posteriorly directed) 30 12. Enp1P3 without seta P. minutus Lang, 1948 29. Dorsal seta attached almost at end of Fu .. Enp1P3 with inner seta 13 ...... P. parafurciger parafurciger Geddes, 1968 13. Exp3P2 and Exp3P3 with 4 setae/spines . Dorsal seta attached at 2/3 of furcal length...... P. tenuis Wells and Rao, 1985 .. P. parafurciger caroliensis Coull, 1971 342 T. Karanovic, G.L. Pesce, W.F. Humphreys

30. Fu with well developed dorsal chitinous ridge. Middle apical seta on Fu long and stout ...... P. furciger Sars, 1907 .. P. ancylus Mielke, 1992 Fu without dorsal ridge 31 46. Exp3P4 with 6 setae/spines . 31. Fu more than 6 times longer than wide ...... P. chavei Coull, 1970 ...... P. curtus Marcus, 1976 Exp3P4 with 7 setae/spines .. Fu about 4 times longer than wide ...... P. galapagoensis Mielke, 1989 ...... P. stigmosus Wells and Rao, 1987 47. Exp3P3 with 5 setae/spines 48 32. Exp2P4 without inner seta 33 Exp3P3 with 6 setae/spines 51 Exp2P4 with inner seta 43 48. A1 with unguiform process on second segment 33. A18-segmented 34 ...... P. parabradyi Lang, 1965 A1 9-segmented 35 A1 without such process 49 49. Abdominal somites with spiniform cuticular 34. Exp3P3 with 4 setae P. danielae Bodin, 1964 extensions P. alatus Fiers, 1986 Exp3P3 with only 3 setae. P. kunzi Mielke, 1989 Abdominal somites without such extensions 50 35. A1 with strong, sharp, unguiform process on 50. ExpP5 with 5 setae in male . second segment 36 ...... P. thiebaudi thiebaudi Petkovski, 1955 A1 without or with small blunt process 37 ExpP5 with 4 setae in male . 36. Exp3P4 with 6 setae and spines ...... P. thiebaudi santacruzensis Mielke, 1989 ...... P. paraborutzkyi Kunz, 1975 51. A1 with strong, sharp, unguiform process on Exp3P4 with 7 setae and spines .. second segment ...... P. carinatus Mielke, 1992 ...... P. setouchinensis Kitazima, 1981 37. A1 with smallblunt process on second segment A1 without or with small blunt process 52 ...... 38 52. A1 without any process on second segment . A1 without any process 39 ...... P. gracilipes Wells and Rao, 1987 38. Exp3P4 with 7 setae/spines, Fu 1.5 times longer A1 with small blunt process 53 than wide P. hibernicus (Roe, 1955) 53. Enp1P1 about as long as ExpP1 . Exp3P4 with 6 setae/spines. Fu 3 times longer ...... P. briani briani Petkovski, 1955 than wide P. hardingi (Roe, 1955) Enp1P1longer than ExpP1 . 39. Exp3P4 with 6 setae/spines ...... P. briani petkovskii Kunz, 1984 stat. novo ...... P. paramossmani (Lang, 1934) Large harpacticoid genera frequently accumulate Exp3P4 with 7 setae/spines 40 numerous synonyms during the course of their 40. Fu about 3.5 times as long as broad .. taxonomic history. The genus Phyllopodopsyllus is ...... P. berrieri Monard, 1936 especially rich with synonyms, many known of long standing. In such a way P. pirgos Apostolov, 1969 is Fu about 1.5 times as long as broad 41 a synonym of P. briani Petkovski, 1955, what was 41. Middle apical seta on Fu not broadened basally noticed by Coull (1973) and accepted by Apostolov ...... P. laspalmensis Marinov, 1973 and Marinov (1988), although Kunz (1984) claimed Middle apical seta on Fu broadened basally ..... it to be a synonym of P. thiebaudi Petkovski, 1955...... 42 Lang (1965) noticed that P. intermedius (Noodt, 1955) is a synonym of P. thiebaudi Petkovski, 1955. 42. ExpP5 with 5 setae in male . Two species were synonymized by their authors ...... P. mossmani T. Scott, 1912 themselves: Kunz (1963) synonymized his P. ExpP5 with 4 setae in male . trichophorus (Kunz, 1951) with P. mossmani T. Scott, ...... P. mossmani chiloensis Mielke, 1992 1912; and Apostolov (1972) synonymized his P. ponticus Apostolov, 1968 with P. pauli Crisafi, 1960. 43. A1 8-segmented 44 P. mielkei mielkei Kunz, 1984 and P. mielkei A1 9-segmented 47 californicus Kunz (1984) are synonyms of P. 44. Exp3P2 and Exp3P3 with 4 setae/spines 45 setouchiensis Kitazima, 1981, which was nicely noticed by Mielke (1992), but also P. crenulatus Exp3P2 with 5, while Exp3P3 with 6 setae/ Wells and Rao, 1987 is a synonym of P. setouchiensis. spines 46 Although Wells and Rao (1987) said that P. 45. Middle apical seta on Fu minute compared to crenulatus "is unique in no single character but the furcal length P. pauli Crisafi, 1960 combination is not matched by any other species of Phyllopodopsyllus wellsi sp. novo from ground waters of Western Australia 343 the genus", furcal rami shape, setal formula, copepods (1997 Edition). Documents traveau l'Institut antennulae, and many other characteristics are quite royal des Sciences naturelles de Belgique, Bruxelles. the same as in P. setouchiensis. P. gertrudi Kunz, 1984 Bradbury, I.H. and Williams, W.D. (1996). Two new and P. gertrudi costaricensis Mielke, 1992 are species of anchialine amphipod (Crustacea: synonyms of P. aegypticus Nicholls, 1944. As a result Hadziidae: Liagoceradocus) from Western Australia. of the description of P. g. costaricensis, Mielke (1992) Records ofthe Western Australian Museum 17: 395-409. unintentionally returned the nominal species P. Bradbury, I.H. and Williams, W.D. (1997). The amphipod gertrudi within the range of the species P. aegypticus, (Crustacea) stygofauna of Australia: description of because the only traditionally used differentiating new taxa (Melitidae, Neoniphargidae, Paramelitidae), character (i.e. distal exopodite segment of third and a synopsis of known species. Records of the Australian Museum 49: 249-34l. swimming leg with 5 or 6 elements) become invalid. Both species have, among other characters, quite the Bruce, N.L. and Humphreys, W.F. (1993). Raptolana pholeta, sp. nov., the first subterranean flabelliferan same extraordinary strong spines on the distal isopod (Cirolanidae) from Australia. endopodite segments of second and third Invertebrate Taxonomy 7: 875-884. swimming legs (see Kunz, 1984; Wells and Rao, Coull, B.e. (1969). Phyllopodopsyllus hermani, a new 1987; and Mielke, 1992). P. yucatanensis Piers, 1995 species of harpacticoid copepod from Bermuda. is an obvious synonym of P. parajurciger Geddes, Crustaceana 16: 27-32. 1968, and it is very strange how Piers (1995) tried to Coull, Re. (1970). Two new species of Phy/lopodopsy/lus make differential diagnosis of his species on the (Copepoda, Harpacticoida) from Bermuda. base of the relative length of some setae. Several Crustaceana 19: 119-124. species are included in the key, because we could Coull, Re. (1973). Harpacticoid copepods (Crustacea) of not claim with the great certainty that they are the family Tetragonicipitidae Lang: a review and synonyms. Thus, the differential character (furcal revision, with keys to the genera and species. length/width ratio) between P. curtus Marcus, 1976 Proceedings ofthe Biological Society ofWashington 86: 9­ and P. stigmosus Wells and Rao, 1987 applied in our 24. key should be used with caution since the Danielopol, D.L., Baltanas, A. and Humphreys, W.F. (in variability of the first species is unknown. Although press). Danielopolina kornickeri sp. n. (Ostracoda: Marcus (1976) mentioned that the furcal index is 9, Thaumatocyprididae) from a western australian from the drawings that she provided it appears that anchialine cave - morphology and evolution. the ratio is not higher than 6.4. P. carinatus Mielke, Zoologica Scripta. 1992 is very similar to P. paraborutzky Kunz, 1975, De Laurentiis, P., Pesce, G.L. and Humphreys, W.F. but the variability of the latter is practically (1999). Copepods from ground waters of Western Australia, IV. Cyclopids from basin and eraton unknown (Kunz, 1975). The same applies to P. aquifers (Crustacea: Copepoda: Cyclopidae). Records galapagoensis Mielke, 1989 and P. chavei Coull, 1970 ofthe Western Australian Museum 19: 243-257. (see Mielke, 1989; Coull, 1970). P. petkovskii Kunz, Fiers, F. (1995). New Tetragonicipitidae (Copepoda, 1984 is so similar to P. briani Petkovski, 1955 that, Harpacticoida) from the Yucatan continental shelf since their populations are located on two (Mexico), including a revision of the genus completely opposite parts of the world, the former Diagoniceps Willey. Bulletin de I'Institut Royal des can be at most considered as a subspecies of P. Sciences Naturelles de Belgique, biologie 65: 151-236. briani, and so has been treated in our key (as P. Geddes, D.e. (1968). Marine biological investigations in briani petkovskii Kunz, 1984 stat. nov.). Two species the Bahamas, 3. Harpacticoid copepods belonging to are not included in the key although they clearly the family Tetragonicipitidae Lang. Sarsia 32: 21-38. belong to the genus Phyllopodopsyllus: P. minor (T. Humphreys, W.F. (1999a). Relict stygofaunas living in and A. Scott, 1903) and P. tristanensis (Wiborg, sea salt, karst and calcrete habitats in arid north­ 1964). These two species are considered incertae western Australia contain many ancient lineages: sedis, because of the incompleteness of their 219-227 in W. Ponder and D. Lunney (eds), The Other descriptions. Lang (1965) and Coull (1973) included 99%. The Conservation and Biodiversity of Invertebrates. P. minor in their keys on the basis of the furcal rami Transactions of the Royal Zoological Society of New shape, but today this is impossible. South Wales, Mosman. Humphreys, W.F. (199%). Physico-chemical profile and energy fixation in Bundera Sinkhole, an anchialine rerniped habitat in north-western Australia. Journal of REFERENCES the Royal Society ofWestern Australia 82: 89-98. Apostolov, A. (1972). Catalogue des Copepodes Humphreys, W.F. (2000). The hypogean fauna of the harpacticoides marins de la Mer Noire. Zoologischer Cape Range peninsula and Barrow Island, north-west Anzeiger 188: 202-254. Australia. In H. Wilkens, D.e. Culver and W.F. Apostolov, A.M. and Marinov, T.M. (1988). 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